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CSIR-UGC (NET)

Life Sciences
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Developmental Biology

1. BASIC CONCEPTS OF DEVELOPMENT POTENCY, COMMITMENT AND SPECIFICATION

Potency is the capacity to differentiate into specialized cell types. In the strictest sense, this
requires stem cells to be either totipotent or pluripotent – to be able to give rise to any mature cell type,
although multipotent or unipotent progenitor cells are sometimes referred to as stem cells.
1. Totipotent Stem cells are produced from the fusion of an egg and sperm cell. Cells
produced by the first few divisions of the fertilized egg are also totipotent. These cells can
differentiate into embryonic and extraembryonic cell types.
2. Pluripotent Stem cells are the descendants of totipotent cells and can differentiate into
cells derived from any of the three germ layers.

Fig. : The Capacity to Differentiate into Specialized Cell Types


3. Multipotent Stem cells can produce only cells of a closely related family of cells (e.g.,
haematopoietic stem cells differentiate into red blood cells, white blood cells, platelets, etc).
4. Unipotent Cells can produce only one cell type, but have the property of self-renewal
Which distinguishes them from non-stem cells (e.g., muscle stem cells).
Differentiation is an overt change in cellular biochemistry; structure, physiology and function. It is
preceded by the covert commitment of cells to particular pre-destined fates. Commitment 18 also a
staged process divided into two stages, specification and determination. Specification implies early,
reversible steps, whereas determination is irreversibility.

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Fig. : An Overview of Developmental Processes


Stages of Cell Commitment
(a) Specification Capable of autonomous differentiation when placed in a neutral environment;
not when placed in non-neutral environment.
(b) Determination Capable of autonomous differentiation even when placed into another
embryonic region.

Fig. : Stages of Cell Commitment : Specification (Reversible) and Determination (Essentially Irreversible)
Specification Types
(a) Autonomous specification
(b) Syncytial specification
(c) Conditional specification
Autonomous Specification
Cells are specified by differential distribution of cytoplasmic components during cleavage of the
egg and early embryo.
Modes of Cell Type Specification and Their Characteristics
 Characteristics of most invertebrates.
 Specification by differential acquisition of certain cytoplasmic molecules present in the
egg.

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 Invariant cleavages produce the same lineages in each embryo of the species. Blastomere
fates are generally invariant.
 Cell type specification precedes any large-scale embryonic cell migration.
 Produces ‘mosaic’ development cells cannot change fate if a blastomere is lost.

Fig. : Autonomous specification in Tunicate (sea squirt). Blastomeres are committed at a very early
stage in mosaic development. If split, each dissociated blastomere pair forms original structures.
Each blastomere contains positional information in the form of specific proteins and genes.

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Syncytial Specification
Nuclear division without cell division, results in cytoplasm with many nuclei.

Fig. : Syncytial Specification Through Morphogen


(A Substance Governing The Pattern of Tissue Development) Gradients.
Conditional Specification
In contrast to the autonomous specification, this type of specification is a cell-extrinsic process
that relies on cues and interactions between cells or from concentration-gradients of morphogens (a
substance governing the pattern of tissue development). Inductive interactions between neighbouring
cells are the most common mode of tissue patterning. In this mechanism, one or two cells from a group
cells with the same developmental potential are exposed to a signal (morphogen) from outside the group.
Modes of Cell Type Specification and Their Characteristics
 Conditional specification.
 Characteristic of all vertebrates and few invertebrates.
 Specification by interactions between cells. Relative positions are important.
 Variable cleavages produce no invariant fate assignments to cells.
 Massive cell rearrangements and migrations precede or accompany specification.
 Capacity for ‘regulative’ development: allows cells to acquire different functions.

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Fig. : Conditional Specification

Fig. : Morphogen Gradients Conditional Specification.


Here, cells respond to protein concentration by turning different colours.
Pattern Formation
Pattern formation is the process by which cells become organized in the embryo, initially to form
the basic rudiments of the body plan and later into the fine structure of individual organs. The process
by which cells become specialized according their position is called regional specification. The first
patterning event that takes place in the embryo is axis specification, the establishment of the principal
body axes (anteroposterior, dorsoventral, etc). These axes may be pre-determined by the distribution of
maternal gene products in the egg. Once the principal axes are established, the position of cells along
each axis is specified and allocating each a positional value. The positional values to cells may be
assigned according to the position of a cell in a morphogen gradient. A diversity of patterning mechanisms
occurs later in development, including asymmetric cell division, specific localized interactions between
cells and the generation of regular spacing patterns by lateral inhibition.
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Induction and Competence


Induction or proximate interaction is the interaction at close range between two or more cells or
tissues with different histories and properties. For example, in the vertebrate eye, the diameter of lens
tissue is controlled by muscle tissue, which meets the tissue of the neural retina. Such coordination in
the construction of organs is accomplished by one group of cells changing the behaviour of an adjacent
set of cells. Not all tissues can respond to the signal being produced by the inducer (tissue that produces
a signal to changes cellular behaviour). For instance, if the optic vesicle (presumptive retina) of Xenopus
laevis is placed in an ectopic location (i.e., in a different place from where it normally forms) underneath
the head ectoderm, it will induce that ectoderm to form lens tissue. Therefore, optic vesicle is an inducer.

Fig. : Ectodermal Ability to Respond to The Optic Vesicle Inducer in Xenopus Laevis
1. The optic vesicle is able to induce lenses in the anterior portion of the ectoderm, but not
in the presumptive trunk and abdomen.
2. If the optic vesicle is removed.
3. The surface ectoderm forms either an abnormal lens or no lens at all.
4. Most other tissues are not able to substitute for the optic vesicle.
Competence is ability to respond to a specific inductive signal. Competence is an actively acquired
condition.

Fig. : Inductive Interactions : Interaction between Epithelia and Mesenchyme : Mesenchyme Plays
An Instructive Role (as The Inducing Tissue); and Initiates Gene Activity in Epithelial Cells.
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Genetic Specificity of Induction


Mesenchyme induces epithelial structures but can only induce what the epithelium is genetically
able to produce.

Fig. : Genetic Specificity showing Epithelial Response is Limited to Genomic Capability


For example, in the developing chick and mammalian eye the Pax6 protein appears to be important
in making the ectoderm competent to respond to the inductive signal from the optic vesicle. Pax6
expression is seen in the head ectoderm, which can respond to the optic vesicle by forming lenses and
it is not seen in other regions of the surface ectoderm.

Fig. : Pax6 is a competence factor for lens induction. During lens induction Pax6 expressed
in the head ectoderm, but not in other regions of surface ectoderm.

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Mesenchymal-Epithelial Interactions
Many inductions involve interactions between epithelia and mesenchyme. Mesenchyme initiates
gene activity in epithelial cells.

Fig. : Epithelial derivatives (hair, scales, feathers, mammary glands, sweat glands)
type depends on restrictions by region and genetics.
In regional specificity, source of the mesenchyme (inducing tissue) determines the structure
of the epithelial derivative.

Fig. : Regional Specificity of Induction


Determination and Differentiation
Cell determination is the process by which portions of the genome are selected for expression
in different embryonic cells. This involves development decisions that gradually restrict cell fate. Cells
can progress from totipotent to pluripotent to determined.

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Molecular Basis of Determination


The result of determination is the presence of tissue-specific proteins characteristic of a cell’s
structure and function.

Fig. : Molecular Basis of Determination of Cells in The Presence of Tissue-Specific Proteins.


In some cases, determination results from the asymmetric segregation of cellular determinants.
However, in most cases, determination is the result of inductive signaling between cells. The sources
of information instruct a cell to express genes at the appropriate time are described below:
(i) Information in the cytoplasm of the unfertilized egg, in the form of RNA and protein, that
is of maternal origin

Fig. : Cytoplasmic Determinants in Egg


(ii) Chemical signals produced by neighbouring embryonic cells, such signals, through a
process called ‘induction’ influence the growth and differentiation of adjacent cells.

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Fig. : Induction of Neighbouring Cells


Cell Differentiation is the process whereby an unspecialized early embryonic cell acquires the
features of a specialized cell such as a heart, liver or muscle cell.
Differentiation follows determination, as the cell elaborates a cell-specific developmental
program. Differentiation results in the presence of cell types that have clear-cut identities, such as
muscle cells, nerve cells, and skin cells.
Differentiated cell remain totipotent as differentiated cells are used in most of the cloning of
animal by nuclear transfer (See table ahead).
Table : Cloning Animals from Differentiated Cells by Nuclear Transfer

Ye ar Animal Nucle us Source


1962 Frog Tadpole intestine
1997 Sheep Mam m ary gland
1998 Cow Adult cum ulus cell
1998 Mouse Adult fibroblast
1999 Goat Foetal fibroblast
2000 Pig Granulosa cell
Principle of Cell Differentiation
(i) Each differentiated cell contains a complete genome.
(ii) The differences between the cell types are only due to expresssion of different kinds of
protein, that is, a set of genes are expressed in a certain type of cell, but not in another
type of cell.
Cell Differentiation is Controlled by Transcription
Tissue Specific transcription factors are the proteins that cause the expression of a group of
proteins that together give cell its identity.
Table : Tissue Specific Transcription Factors That Cause Cell Differentiation into Specific Tissue

Organism Factor Type of Factor Tissue Specified


Mouse SCL/Ta/1 Basic helix-loop-helix Haematopoetic cells
Frog (Xenopus) neuroD Basic helix-loop-helix Neurons
Mouse Tcf4 HMG box Intestinal epithelium
Mouse MyoD Basic helix-loop-helix Skeletal muscle
Mouse Pax7 Paired-box Skeletal muscle

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Fig. : Muscle Differentiation, Transcription Factors in Muscle Differentiation


Cell differentiation is reversible :
(a) Embryonic Stem (ES) cell can complete the whole process of differentiation in proper
environment.
(b) Differentiated cell can undergo dedifferentiation or trans-differentiation.
(c) Cancer cell can be induce to differentiate.
Morphogenetic Gradients
Morphogenesis is the process of cellular differentiation, distribution and growth that takes place
during the embryonic development of an organism. The morphogenes (proteins that control
morphogenesis) that determine the fate of cells and elicits different responses at different concentration
thresholds. A morphogen synthesized at one end of an axis would establish a concentration gradient
along the axis. The gradient of morphogen is translated into a gradient of intracellular signal transduction
and ultimately differential gene expression. Morphogens can establish gradients by free diffusion but
alternative mechanisms are required for extracellular secreted proteins such as Hedgehog, Wingless/
Wnt and Decapentaplegic/BMP
The morphogenetic determinants (certain proteins or mRNAs) are placed in different regions of
the egg cytoplasm and are apportioned to the different cells as the embryo divides. These morphogenetic
determinants specify the cell type.
Morphogenetic Gradients in Drosophila Melenogaster
The morphogenes (or transcription factors) displayed are the proteins bicoid (bicoid is a maternally
transcribed gene that organizes the anterior development in Drosophila) and hunchback (hunchback is
the ‘partner’ of bicoid in anterior/posterior development in Drosophila). These proteins play an important
role in determining the anterior part of the body (head and thorax) from the posterior part (abdomen).
In Drosophila, antennae and legs are created by the same process, the only differ is a single
transcription factor. If this transcription factor is damaged, the fly grows legs instead of antennae. This
is due to ‘antennapedia’ mutant.
In the early stages of morphogenesis in an insect embryo, four types of differentiation can be
distinguished :
 Anterior (head and thorax)
 Posterior (abdomen)

Fig. : Schematic Drawing of Morphogene Gradients in Drosophila


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(a) The bicoid mRNA (messenger RNA) is an intermediate copy of a piece of the DNA. It is
transported to the ribosomes, where protein biosynthesis takes place) is located on the
left side of the embryo. All cells of the embryo have inactive hunchback DNA.
(b) The bicoid mRNA is expressed as bicoid protein, resulting in a protein gradient with the
highest concentration of protein on the left side of the embryo.
(c) The hunchback DNA is activated once the amount of bicoid protein passes a certain
threshold. This results in a sharp borderline, which divides the part, where hunchback is
expressed from the part where hunchback is not expressed.
 Dorso-ventral (front-and backside)
 Terminal (special structures at the unsegmented ends of the embryo)
In the later stages of development, these basic compartments become more and more divided.
The same transcription factor can be used several times, having different meanings in the different body
parts.
The term ‘morphogenesis’ can be used to describe the development of unicellular life forms that
do not have an embryonic stage in their life cycle. Morphogenetic responses may be induced in organisms
by hormones or by environmental chemicals.
Cell Fate and Cell Lineages
Fate is the sum of all structures that the cell or its descendants will form at a later stage of normal
development.
It is the ultimate differentiated state to which a cell has become committed. Fate maps are the
bases for experimental embryology
The cell lineage of an organism is the pattern of cell divisions during its development. Cell
lineages are described by following cell divisions in living individuals or by marking cells and examining
their progeny.
Cell Lineage Patterns
(i) Some organisms or precursor cells display cell lineage patterns are invariant patterns of
cell division, in which specification of cell fates is correlated with cell division patterns.
(ii) In other organisms, cell lineage patterns are variable and not correlated with cell fates.
Invariant cell lineages reflect both cell-autonomous mechanisms of fate determination and
highly reproducible cell-cell interactions. Genetic analysis of cell lineages has focused on
systems, where cell lineage and cell fates are correlated.
Fate Mapping Methods
Fate maps have been generated in several ways:
(i) Observing living embryos In certain invertebrates, it is actually possible to look through
the microscope and trace the descendants of a particular cell into the organs they generate.
This type of study was performed about a century ago by Edwin G Conklin.

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Fig. : Fate of Three Germ Layers


(b) Vital dye marking In the early years of the twentieth century, Vogt (1929) traced the fates
of different areas of amphibian eggs by applying vital dyes to the region of interest. Vital
dyes will stain cells but not kill them.

Fig. : Vital dye staining of amphibian embryos. (a) Specific cells of the embryonic surface with vital dyes.
(b-d) Dorsal surface views of stain on successively later embryos. (e) Medial sagittal section to
show the stained cells
(iii) Radioactive labelling and fluorescent dyes In this method, a donor embryo is usually
grown in a solution containing radioactive thymidine. This base will be incorporated into
the DNA of the dividing embryo.
The problems with radioactive and vital dye marking include their dilution over many cell divisions
and the laborious preparation of the slides.
Methods of Analyses of Cell Lineage
(i) Direct observation Same as described above under ‘Fate Mapping Methods’.
(ii) Clonal analysis In large, opaque or slowly developing embryos, direct observation of cell
divisions is not feasible. To analyze cell lineages in such cases, it is necessary to mark
individual cells by physical or genetic means and later to identify their progeny by expression
of the marker. Here, the progeny of a single cell forms a clone.
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Stem Cells
Stem cells are a class of undifferentiated cells (i.e., lacking certain tissue specific differentiation
markers) that are able to differentiate into specialized cell types.
All stem cells have three general properties.
1. Stem cells are capable of dividing and renewing themselves for long periods.
2. Stem cells are unspecialized.
3. Stem cells can give rise to specialized cells.
The two broad types of mammalian stem cells are embryonic stem cells that are found
in blastocysts and adult stem cells that are found in adult tissues.
In a developing embryo, stem cells can differentiate into all of the specialized embryonic
tissues.
4. Embryonic stem cells are primitive (undifferentiated) cells from the embryo that have the
potential to become a wide variety of specialized cell types.
5. Human embryonic stem cells are derived from blastocyst.
Embryonic stem cells develop from eggs that have been fertilized invitro or invivo. Another
potential source of embryonic stem cells is Somatic Cell Nuclear Transfer (SCNT). Adult
stem cell is an undifferentiated cell found among differentiated cells in a tissue or organ.
One population is haematopoietic stem cells and the second population is bone marrow
stromal cells.
Genomic Equivalence and The Cytoplasmic Determinants
Genomic equivalence means that the genornes of all or most of the somatic cells within a
multicellular organism are essentially the same. Thus, the differentiation of cells does not (usually)
involve changes in the nucleotide sequence of DNA. Genomic equivalence can be demonstrated via
various cloning experiments including the growing of a whole plant from an isolated cell (other than a
zygote) and the cloning of an animal via the injection of a somatic-cell genome into a zygot (or unfertilized
egg) whose own genome has been destroyed.
Genomic Equivalence for Plants and Animals
 Genomic equivalence; nearly all of the cells of an organism have the same genes.
 Because the cells of animals will not often divide in culture, scientists have adopted
alternative approaches to examine genomic equivalence: transplanting nuclei of differentiated
cells into enucleated egg cells of frogs.
 Plants’ genomic equivalence was demonstrated by experiments in which entire individuals
developed from differentiated somatic cells.

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Evidence for Genomic Equivalence

Fig. : Procedure for transplanting blastula nuclei into activated enucleated Rana pipiens eggs.
The relative dimensions of the meiotic spindle have been exaggerated to show the technique

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Fig. : Nuclei of Adult Somatic Cells are used for Cloning Mammals
(a) The upper left dark mouse is the oocyte donor, while the upper right light (agouti) mouse
is the nucleus donor. The white mouse in the lower row is the mouse into whose uterus
the resulting embryos were implanted. The two agouti mice beside her are cloned mice,
derived from the injection of the agouti nucleus into the oocyte of the black-furred parent.
(b) Similarly, Dolly, the adult sheep, was derived by fusing a mammary gland cell nucleus with
an enucleated oocyte, which was then implanted in a surrogate mother (of a different
breed of sheep who gave birth to dolly.)
Table : Examples of Genomic Non-equivalence During Development

Mechanism Examples
DNA amplification Selective amplificaton of rRNA genes in Xenopus oocytes.
Amplification of the chorion genes during oogenesis in
Drosophila.
DNA loss In mammals, loss of nuclei in differentiated erythrocytes
and keratinocytes.
DNA rearrangement In vertebrates, VDJ recombination and class switching
during B-cell and T-cell development.
A cytoplasmic determinant is a substance or substances, located in part of an egg or blastomere,
that guarantees the assumption of a particular state of commitment by the cells which inherit it during
cleavage.
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Cytoplasmic determinants are of considerable importance for the very earliest stages of embryonic
development because they are often responsible for the establishment of the first two or three distinctly
specified regions in the embryo. There are established experimental techniques that reorganize the
cytoplasm and thus redistribute the determinants or block their transport. For example, the transfer of
cytoplasm from the anterior pole to the posterior of the Drosophila egg by microinjection demonstrates
the role of polarized maternal cytoplasmic determinants in axis specification, since the embryo develops
with two heads and no abdomen.
In the early stages of C. elegans development a complex containing PAR3, PAR6 and aPKC
becomes localized in the anterior and controls the fate of the first two blastomeres. A similar system
seems to be involved in later events of neurogenesis and epithelial polarization in other animals.
Imprinting
Genomic imprinting is an epigenetic phenomenon which, in most cases, is believed to occur in
gametogenesis. Genomic imprinting occurs when both maternal and paternal alleles are present, but one
allele will be expressed, while the other remains inactive. It is not completely evident why genes are
imprinted. The most prominent assumption is that this process is necessary for development and may
somehow regulate growth in the embryo and neonate. Evidence for this suggestion came from experiments
with androgenotes (embryos with two paternal genomes) and gynogenotes (embryos with two maternal
genomes), which were produced by nuclear transplantation. These zygotes were formed, but neither
type was able to undergo further development. From this situation, it is possible to suggest that the
maternal and paternal effects are complementary.
The optimal method for gene imprinting is DNA methylation. This process is carried out with the
enzyme DNA methyltransferase (DNA MTase) in mammals. DNA MTase acts on the DNA sequence 5-
CpG-3. Some organisms (primarily higher eukaryotes) have aggregates of CpG (known as CpG islands)
in their genomes. These islands are rarely methylated in animal cells. This may be due to the bound
transcription factors that block DNA MTase.
Mutants and Transgenics in Analysis of Development
Recently developed techniques have enabled us to study gene function by moving certain genes
into and out of embryonic cells.
Technique of Inserting DNA into a Cell
Cloned pieces of DNA can be isolated, modified (if so desired) and inserted into cells by several
means. Microinjection, in which a solution containing the cloned gene is injected into the nucleus of a
cell. This technique is especially useful for inserting genes into newly fertilized eggs, since the eggs are
relatively large. Transfection, DNA fragments may be incorporated directly into cells by incubating the
cells in a solution containing new DNA. The chances of a DNA fragment incorporation into the chromosomes
are relatively small. Electroporation, in which a high-voltage pulse ‘pushes’ the DNA into the cells.
Transposable element or retroviral vector, a more ‘natural’ way of getting genes into cells is to insert
the cloned gene into a transposable element or retroviral vector. These naturally occurring mobile regions
of DNA can integrate themselves into the genome of an organism.
Chimeric Mice Experiments
In the blastocyst, there is a stage when two cell types are present: The outer trophoblast cells,
which will form the fetal portion of the placenta and the Inner Cell Mass (ICM), whose cells will give rise
to the embryo itself. Separation of ICM cells can lead to twins and if an 1CM blastomere from one mouse
is transferred into the embryo of a second mouse, the donor 1CM cell can contribute to every organ of
the host embryo.
Inner cell mass blastomeres can be isolated from an embryo and cultured in vitro, such cultured
cells are called embryonic stem cells (ES cells). ES cells are almost totipotent, except the trophoblast.
Embryonic stem cells from a mouse are cultured and their genome altered by the addition of a
cloned gene. These transgenic cells are selected and then injected into the early stages of a host mouse
embryo.
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The transgenic embryonic stem cells integrate with the host’s embryonic stem cells. The embryo
is placed into the uterus of a pregnant mouse, where it develops into a chimeric mouse. The treated
cells can become part of the germ line of the mouse. If such a chimeric mouse is mated with a wild-
type mouse, some of its progeny will carry one copy of the inserted gene. When these heterozygous
progeny are mated to one another, about 25 percent of the resulting offspring will be homozygous for
the inserted gene in every cell of their bodies. Thus, a gene cloned from some other organism will be
present in both copies of the chromosomes within these mouse genomes. Strains of such transgenic
mice have been particularly useful in determining how genes are regulated during development.

Fig. : Production of Transgenic Mice


Gene Targeting (Gene Knockout) Experiments
The analysis of early mammalian embryos has been done by the techniques of gene targeting
(gene knockout). This technique involves adding genes, gene targeting replaces wild-type alleles with
mutant ones. As an example, a knockout of the gene for bone morphogenetic protein 7 (Bmp7) is
considered in the following discussion.
Bone morphogenetic proteins are involved in numerous developmental interactions. Bmp7 has
been implicated as a protein that prevents cell death and promotes cell division in several developing
organs.

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To find the function of Bmp7 in the development of the mouse. First, isolate the (Bmp7) gene,
cut it at one site with a restriction enzyme and inserted a bacterial gene for neomycin resistance into
that site. This mutated the Bmp7 gene has destroyed the ability of the gene’s product (Bmp7 protein)
to function. These mutant (Bmp7) genes were electroporated into embryonic stem cells that were
naturally sensitive to neomycin. Once inside the nucleus of an ES cell, the mutated Bmp7 gene may
replace a normal allele of Bmp7 by a process called homologous recombination. Thus, the enzymes
involved in DNA repair and replication incorporate the mutant gene in the place of the normal copy. It’s
a rare event, but such cells can be selected by growing the ES cells in neomycin. The resulting cells
have one normal Bmp7 gene and one mutated Bmp7 gene. These heterozygous ES cells were then
microinjected into mouse blastocysts, where they were integrated into the cells of the embryo. The
resulting chimeric mice had wild-type cells from the host embryo and heterozygous Bmp 7-containing
cells from the donor ES cells. The chimeras were mated to wild-type mice, producing progeny that were
heterozygous for Bmp7. These heterozygous mice were then bred with each other and about 25 percent
of their progeny carried two copies of the mutated Bmp7 gene.
The homozygous mutants lacked eyes and kidneys. In the absence of functional Bmp7 protein,
it appears that many of the cells that normally form these two organs stop dividing and die. In this way,
gene targeting can be used to analyze the roles of particular genes during mammalian development.

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Fig. : Technique for Gene Targeting. In this case, the targeted gene is Bmp7
(a) Embryonic stem cells from a mouse blastocyst are cultured.
(b) Cloned Bmp7 genes are cut with a restriction enzyme and a neomycin resistance gene
is inserted into the region that encodes the DNA-binding site of the protein. These mutant
Bmp7 genes are electroporated into ES cells. In some of the cells, homologous
recombination exchanges a wild-type gene for the mutant copy. These cells are selected
by their neomycin resistance.

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Question with Solutions

1. In higher plants, the red/far-red sensory photoreceptor, phytochrome, is a light regulated kinase.
Which of the following classes of kinase does it represent ?
(A) Two-component sensor regulator (histidine kinase)
(B) Two-component sensor regulator (serine/threonine kinase)
(C) Reprogramming of the shoot apical meristem
(D) Synthesis of the flowering hormone florigen

2. Match the List I and List II


List – I List – II
Stages of egg maturation at the Animal Species
time of sperm entry
1. Primary oocyte A. Birds
2. First Metaphase B. Most mammals
3. Second Metaphase C. Many insects
4. Meiosis Complete D. Sea Urchin
Codes :
1 2 3 4
(A) A C B D
(B) A B C D
(C) A C D B
(D) C A D B

3. In terms of definition for stem cells, a fertilized egg could be considered as


(A) Omnipotent (B) Multipotent
(C) Pluripotent (D) Totipotent

4. What does not happen during early gastrutation ?


(A) Three embryonic germ layers are established
(B) Large scale migration of cells to organise three germ layers
(C) The embryo increase in size
(D) Development fate of germ layers are decided.
5. Homeotic gene sequence
(A) Are found from fruit to human beings
(B) Control segmentation and basic patterns of development of body parts
(C) Are very different but present among nearly all eukaryotes
(D) Only A & B

6. In human female somatic cells :


(A) The paternally derived X chromosome is preferentially inactivated.
(B) The maternally derived X chromosome is preferentially inactivated.
(C) One of the X chromosome is inactivated randomly in cells.
(D) One of the X chromosome is lost.

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Life Sciences-LS (Sample)

7. A Graffian follicle is
(A) An immature developing follicle (B) A mature follicle ready to ovulate
(C) A follicle undergoing apoptosis (D) An ovulated follicle

8. The diagram represents the top view of a developing egg at 8 celled stage of cleavage. Which
one of the following group of animals show this pattern

(A) Platyhelminthes (B) Echinodermata


(C) Ctenophora (D) Cnidaria

9. An embryo lacking bicoid is injected with bicoid m-RNA at middle portion. It will result into
(A) Two heads and no telson
(B) Head in the middle and telson at both ends
(C) No head and telson at both ends
(D) Normal phenotype

10. Which one is not associated with gametogenesis?


(A) Formation of ova (B) Formation of spermatid
(C) Release of ova (D) Change of spermatids of spermatozoa

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Life Sciences-LS (Sample)

SOLUTIONS

1. (B) Phytochromes may function as light-regulated serine/threonine kinases, and can


phosphorylate several substrates, including themselves in vitro.
2. (A) List – I List – II
Stages of egg maturation at the Animal Species
time of sperm entry
1. Primary oocyte A. Birds
2. First Metaphase C. Many insects
3. Second Metaphase B. Most mammals
4. Meiosis Complete D. Sea Urchin
3. (D) Totipotent stem cells are perhaps the most versatile of the stem cell types. As explained,
a totipotent zygote cell is created when a single celled sperm and egg unite. This totipotent
fertilised egg has the potential to give rise to virtually all human cells, such as nerve or
heart.
4. (C) The embryo increase in size
5. (D) Homeoboxes have been found in many organisms from fruit flies to human. Homeotic
gene, any of a group of genes that control the pattern of body formation during early
embryonic development of organisms. These genes encode proteins called transcription
factors that direct cells to form various parts of the body.
6. (C) One of the X chromosome is inactivated randomly in cells.
7. (B) A fluid-filled structure in the mammalian ovary within which an ovum develops prior to
ovulation.
8. (A) The simplest animals that are bilaterally symmetrical and triploblastic (composed of three
fundamental cell layers) are the Platyhelminthes, the flatworms. Flatworms have no body
cavity other than the gut (and the smallest free-living forms may even lack that!) and lack
an anus; the same pharyngeal opening both takes in food and expels waste. Because of
the lack of any other body cavity, in larger flatworms the gut is often very highly branched
in order to transport food to all parts of the body.
9. (B) Head in the middle and telson at both ends
10. (C) Release of ova

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